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Biological control 1 Biological

The current scope of article does not include composting techniques. For these see Composting: Destroying , , or unwanted or Mulch: Mulching (composting) over unwanted plants. Biological control is a method of controlling pests (including , , and ) using other living . It relies on , , herbivory, or other natural mechanisms, but typically also involves an active management role. It can be an important component of integrated pest management (IPM) programs. There are three basic types of strategies: importation (sometimes called classical biological control), augmentation and conservation. Natural enemies of pests, also known as biological control agents, include predators, , and pathogens. Biological control agents of plant diseases are most often referred to as antagonists. Biological control agents of weeds include and plant pathogens.

Types of biological pest control There are three basic types of biological pest control strategies: importation (sometimes called classical biological control), augmentation and conservation.[1]

Importation Importation (or "classical biological control") involves the introduction of a pest's natural enemies to a new locale where they do not occur naturally. This is usually done by government authorities. In many instances the complex of natural enemies associated with a pest may be inadequate, a situation that can occur when a pest is accidentally introduced into a new geographic area, without its associated natural enemies. These introduced pests are referred to as exotic pests and comprise about 40% of the insect pests in the . The process of importation involves determining the origin of the introduced pest and then collecting appropriate natural enemies associated with the pest or closely related . Selected natural enemies are then passed through a rigorous assessment, testing and process, to ensure that they will work and that no unwanted organisms (such as hyperparasitoids) are introduced. If these procedures are passed, the selected natural enemies are mass produced and then released. Follow-up studies are conducted to determine if the natural enemy becomes successfully established at the site of release, and to assess the long-term benefit of its presence. To be most effective at controlling a pest, a biological control agent requires a colonizing ability which will allow it to keep pace with the spatial and temporal disruption of the . Its control of the pest will also be greatest if it has temporal persistence, so that it can maintain its population even in the temporary absence of the target species, and if it is an opportunistic forager, enabling it to rapidly exploit a pest population.[2] However an agent with such attributes is likely to be non- specific, which is not ideal when considering its overall ecological impact, as it may have unintended effects on non-target organisms. There are many examples of successful importation programs, including: • Joseph Needham noted a Chinese text dating from 304AD, Records of the Plants and Trees of the Southern Regions, by Hsi Han, which describes mandarin oranges protected by biological pest control techniques that are still in use today. • One of the earliest successes in the west was in controlling purchasi, the cottony cushion scale, a pest that was devastating the citrus industry in the late 19th century. A predatory insect Rodolia cardinalis (the Vedalia ), and a were introduced from by Charles Valentine Riley. Within a few years the cottony cushion scale was completely controlled by these introduced natural enemies. • Damage from postica Gyllenhal, the , a serious introduced pest of forage, was substantially reduced by the introduction of several natural enemies. 20 years after their introduction the population of Biological pest control 2

in the alfalfa area treated for alfalfa weevil in the Northeastern United States was reduced by 75 percent. • A small , ostriniae, was introduced from to help control the ( nubilalis), one of the most destructive insects in , making it a recent example of a long history of classical biological control efforts for this pest. • The population of Levuana irridescens (the Levuana ), a serious pest in , was brought under control by a classical biological control program in the 1920s. Classical biological control is long lasting and inexpensive. Other than the initial costs of collection, importation, and rearing, little expense is incurred. When a natural enemy is successfully established it rarely requires additional input and it continues to kill the pest with no direct help from and at no cost. However importation does not always work. It is usually most effective against exotic pests and less so against native insect pests. The reasons for failure are not often known but may include the release of too few individuals, poor of the natural enemy to environmental conditions at the release location, and lack of synchrony between the life of the natural enemy and host pest.

Augmentation Augmentation involves the supplemental release of natural enemies, boosting the naturally occurring population. Relatively few natural enemies may be released at a critical time of the season (inoculative release) or millions may be released (inundative release). An example of inoculative release occurs in production of several crops. Periodic releases of the parasitoid, formosa, are used to control greenhouse , and the predatory Phytoseiulus persimilis is used for control of the two-spotted mite. Lady , lacewings, or parasitoids such as those from the Trichogramma are frequently released in large numbers (inundative release). Recommended release rates for Trichogramma in vegetable or field crops range from 5,000 to 200,000 per acre (1 to 50 per square metre) per week depending on level of pest . Similarly, entomopathogenic are released at rates of millions and even billions per acre for control of certain -dwelling insect pests. The spraying of analogues (such as 3-FMC) has been suggested as a way to boost the effectiveness of augmentation. Octopamine, regarded as the counterpart of plays a role in activating the insects' -or-fight response. The behind using octopamine analogues to augment biological control is that natural enemies will be more effective in their eradication of the pest, since the pest will be behaving in an unnatural way because its flight-or-fight mechanism has been activated. Octopamine analogues are purported to have two desirable characteristics for this type of application: (1) they affect insects at very low dosages (2) they do not have a physiological effect in humans (or other ).[3] Biological pest control 3

Conservation The conservation of existing natural enemies in an environment is the third method of biological pest control. Natural enemies are already adapted to the habitat and to the target pest, and their conservation can be simple and cost-effective. Lacewings, lady beetles, hover fly larvae, and parasitized mummies are almost always present in aphid colonies. Cropping systems can be modified to favor the natural enemies, a practice sometimes referred to as habitat manipulation. Providing a suitable habitat, such as a shelterbelt, hedgerow, or beetle bank where beneficial insects can live and reproduce, can help ensure the survival of populations of natural enemies. -rich plants that bloom for long periods are especially good, as many beneficials are nectivorous during the adult stage, but parasitic or predatory as larvae. A good example of this is the which is frequently found on as an adult, but whose larvae eat , , , and other beetles. In California prune trees are sometimes planted in vineyards to provide an improved habitat or for a key grape pest parasitoid. The prune trees harbor an alternate host for the parasitoid, which could previously overwinter only at great distances from most vineyards. A turnaround , filled with to The provisioning of artificial shelters in the form of wooden caskets, attract Dermaptera-species boxes or is sometimes undertaken, particularly in , to make a cropped area more attractive to natural enemies. For example, the stimulation of the natural predator Dermaptera is done in gardens by hanging upside-down flowerpots filled with straw or wool. Conservation strategies such as mixed plantings and the provision of flowering borders can be more difficult to accommodate in large-scale production. There may also be some conflict with pest control for the large producer, because of the difficulty of targeting the pest species, also refuges may be utilised by the pest insects as well as by natural enemies. Some plants that are attractive to natural enemies may also be hosts for certain plant diseases, especially plant that could be vectored by insect pests to the crop. Naturally occurring biological controls are often susceptible to the same used to target their pest hosts. Biological pest control 4

Biological control agents

Predators

Predators are mainly free-living species that directly consume a large number of prey during their whole lifetime. Ladybugs, and in particular their larvae which are active between May and July in the northern hemisphere, are voracious predators of aphids, and will also consume mites, scale insects and small caterpillars. The larvae of many species principally feed upon greenfly, one devouring up to fifty a day, or 1000 in its lifetime. They also eat fruit tree spider mites and small caterpillars. Adults feed on nectar and , which they require for production.

Lacewings are available from biocontrol dealers.

Dragonflies are important predators of mosquitoes, both in the water, where the naiads eat larvae, and in the air, where adult capture and eat adult mosquitoes. Community-wide programs that spray adult mosquitoes also kill dragonflies, thus reducing an important biocontrol agent. Phasmarhabditis hermaphrodita is a microscopic that kills , thereafter feeding and reproducing inside. The nematode is applied by watering onto moist soil, and gives protection for up to six weeks in optimum conditions. Other useful predators include lacewings, pirate bugs, rove and ground beetles, aphid , , , predatory mites, as Predatory wasp looking for bollworms or other caterpillars on a cotton plant well as larger such as , toads, , , slow- and . Cats and terriers kill field mice, , June bugs, and birds. Dachshunds are bred specifically to fit inside tunnels underground to kill . More examples: • Phytoseiulus persimilis (against spider mites) • Amblyseius californicus (against spider mites) • Amblyseius cucumeris (against spider mites)[4] • (against spider mites, , and white ) • (against spider mites) • punctillum (against spider mites) • caluginosus (against spider mites) Biological pest control 5

Parasitoid insects Parasitoids lay their eggs on or in the of an insect host, which is then used as a for developing larvae. The host is ultimately killed. Most insect parasitoids are or flies, and usually have a very narrow host range. Four of the most important groups are: • Ichneumonid wasps: (5–10 mm). Prey mainly on caterpillars of and . • Braconid wasps: Tiny wasps (up to 5 mm) attack caterpillars and a wide range of other insects including greenfly. A common parasite of the white - seen as clusters of sulphur yellow cocoons bursting from collapsed caterpillar skin. • Chalcid wasps: Among the smallest of insects (<3 mm). Parasitize eggs/larvae of greenfly, whitefly, cabbage caterpillars, scale insects and Moth ( comariana). • Tachinid flies: Parasitize a wide range of insects including caterpillars, adult and larval beetles, true bugs, and others. Examples of parasitoids: • A small predatory which is a parasitoid of whitefly, a sap-feeding insect which can cause wilting and black sooty moulds. It is most effective when dealing with low level , giving protection over a long period of time. The wasp lays its eggs in young whitefly 'scales', turning them black as the parasite larvae pupates. Diagram illustrating the life cycles of and its • Eretmocerus spp. (against white Encarsia formosa flies)[5] • colemani (against aphids) • ashmeadi (: Mymaridae) has been introduced to control the glassy-winged vitripennis (Hemipterae: Cicadellidae) in French and has successfully controlled ~95% of the pest density.[6] Parasitoids are one of the most widely used biological control agents. Commercially there are two types of rearing systems: short-term daily output with high production of parasitoids per day, and long-term low daily output with a range in production of 4-1000million female parasitoids per week.[7] Larger production facilities produce on a yearlong basis, whereas some facilities will produce only seasonally. Rearing facilities are usually a significant distance from where the agents will be used in the field, and transporting the parasitoids from the point of production to the point of use can pose problems. Shipping conditions can be too hot, and even vibrations from planes or trucks can disrupt the parasitoids.[7] Biological pest control 6

Micro-organisms Pathogenic micro-organisms include , fungi, and viruses. They kill or debilitate their host and are relatively host-specific. Various microbial insect diseases occur naturally, but may also be used as biological pesticides. When naturally occurring, these outbreaks are density-dependent in that they generally only occur as insect populations become denser.

Bacteria Bacteria used for biological control infect insects via their digestive tracts, so insects with sucking parts like aphids and scale insects are difficult to control with bacterial biological control.[8] thuringiensis is the most widely applied species of bacteria used for biological control, with at least four sub-species used to control Lepidopteran (moth, ), Coleopteran (beetle) and Dipteran (true flies) insect pests. The bacteria is available in sachets of dried spores which are mixed with water and sprayed onto vulnerable plants such as and fruit trees.

Fungi Fungi that cause in insects are known as entomopathogenic fungi, including at least fourteen species that attack aphids.[9] bassiana is used to manage a wide variety of insect pests including: , thrips, aphids and weevils. A remarkable additional feature of some fungi is their effect on plant . species may enhance biomass production promoting development, dissolving insoluble phosphate containing minerals. Examples of entomopathogenic fungi: • (against white flies, thrips, aphids and weevils) • Paecilomyces fumosoroseus (against white flies, thrips and aphids) • spp. (against beetles, and , , spider mites and other pests) • spp. (against white flies, thrips and aphids) • species (includes teleomorphs of the above genera: that infect a wide spectrum of ) • Trichoderma species are used to manage certain plant pathogens. has been used against , and to treat the spread of fungal and bacterial growth on tree wounds. It may also have potential as a means of combating silver disease.

Viruses

• A viral biological control which can be introduced in to control the of in Australia is the rabbit haemorrhagic disease that causes the rabbit haemorrhagic disease.

Combined use of parasitoids and pathogens

In cases of massive and severe infection of invasive pests, techniques of pest control are often used in combination. An example being, that of the emerald ( planipennis Fairmaire, ), an invasive beetle from China, which has destroyed tens of millions of ash trees in its introduced range in North America. As The European Rabbit (Oryctolagus cuniculus) is part of the campaign against the (EAB), American seen as a major pest in Australia and New scientists in conjunction with the Chinese Academy of Zealand. Biological pest control 7

searched since 2003 for its natural enemies in the wild leading to the discovery of several parasitoid wasps, namely planipennisi, a gregarious larval endoparasitoid, agrili, a solitary, parthenogenic egg parasitoid, and agrili, a gregarious larval ectoparasitoid. These have been introduced and released into the United States of America as a possible biological control of the emerald ash borer. Initial results have shown promise with Tetrastichus planipennisi and it is now being released along with Beauveria bassiana, a fungal with known insecticidal properties.[10][11][12]

Plants The vine pruriens is used in the countries of and as a biological control for problematic grass. is said not to be invasive outside its cultivated area.[13] uncinatum can be used in push-pull farming to stop the , .[14]

Indirect control Pests may be controlled by biological control agents that do not prey directly upon them. For example the Australian bush fly, vetustissima, is a major nuisance pest in Australia, but native decomposers found in Australia are not adapted to feeding on cow dung, which is where bush flies breed. Therefore the Australian Project (1965-1985,) led by Dr. George Bornemissza of the Commonwealth Scientific and Industrial Research Organisation, released forty-nine species of dung beetle,[15] with the aim of reducing the amount of dung and therefore also breeding sites of the fly.[16]

Effects of biological control

Effects on native

Biological control can potentially have positive and negative effects on biodiversity. [2] The most common problems with biological control occur via predation, parasitism, pathogenicity, , or other attacks on non-target species. [17] Often a biological control agent is imported into an area to reduce the competitive advantage of an exotic species that has previously invaded or been introduced there, the aim being to thereby protect the existing native species and ecology. However the introduced control does not always target only the intended species; it can also target native species.[18] In during The , Bufo marinus the 1940s parasitic wasps were introduced to control a lepidopteran pest and the wasps are still found there today. This may have a negative impact on the native , however, host range and impacts need to be studied before declaring their impact on the environment. [19]

Over the past 15 years with the rise in biological control interest there has become a greater focus on the non-target impacts that could occur. [2] In the past many biological control releases were not thoroughly examined and agents of biological control were released without any consideration. When introducing a biological control agent to a new area, a primary concern is its host-specificity. Generalist feeders (control agents that are not restricted to preying on a single species or a small range of species) often make poor biological control agents, and may become themselves. For this reason potential biological control agents should be subject to extensive testing and quarantine before release into any new environment. If a species is introduced and attacks a native species, the biodiversity in that area can change dramatically. When one native species is removed from an area, it may have filled an essential . When this niche is absent it may directly affect the entire ecosystem. Biological pest control 8

Vertebrate tend to be generalist feeders, and seldom make good biological control agents; many of the classic cases of "biocontrol gone awry" involve vertebrates. For example the cane toad, Bufo marinus, was intentionally introduced to Australia to control the introduced French’s Cane Beetle and the Greyback Cane Beetle,[20] pests of sugar cane. 102 toads were obtained from Hawaii and bred in captivity to increase their numbers until they were released into the sugar cane fields of the tropic north in 1935. It was later discovered that the toads could not jump very high and so they could not eat the cane beetles which stayed up on the upper stalks of the cane plants. However the toad thrived by feeding on other insects and it soon spread very rapidly; it took over native habitat and brought foreign disease to native toads and frogs, dramatically reducing their populations. Also when it is threatened or handled, the cane toad releases from parotid glands on its shoulders; native Australian species such as goannas, tiger , and northern quolls that attempted to eat the toad were harmed or killed.[21] This example shows how small mis-introduced organisms can alter the native biodiversity in large . If native species are reduced or eradicated, a domino effect can take place until a new equilibrium is reached. Other examples of biological control agents that subsequently crossed over to native species are: • conicus, a -feeding weevil, was introduced to North America to control exotic thistles (Musk and Canadian). However the weevil does not target only the exotic thistles; it also targets native thistles that are essential to various native insects which rely solely on native thistles and do not adapt to other plant species. • The was introduced to Hawaii in order to control the rat population. However it preyed on the endemic birds of Hawaii, especially their eggs, more often than it ate the rats. (Note, however, that the introduction of the mongoose was not undertaken based on scientific—or perhaps any—understanding of the consequences of such an action. The introduction of a generalist for biocontrol of anything would be unthinkable by any reasonable standards today.) • 5 cats brought to the subantarctic Marion Islands to catch mice in 1949 multiplied to 3,400 in about two decades and started to threaten local of birds. They had to be infected with feline distemper virus. The rest were shot and completely eliminated by the 1990s. • The sturdy and prolific mosquito fish was introduced from around the Gulf of to around the world in the 1930s and 40s to combat ; however, it was found to cause the decline of local fish and frogs through competition of other food source as well as their eggs.[22] (See Mosquitofish in Australia) Living organisms, through the process of , may achieve increased resistance to biological, chemical, and physical methods of control over time. In the event the target pest population is not completely exterminated or is still capable of reproduction (were the pest control means a form of sterilization), the surviving population could acquire a tolerance to the applied pressures - this can result in an evolutionary arms race with the control method. Successful biological control reduces the population density of the target species over several years, thus providing the potential for native species to re-establish. In addition, and reestablishment programs can aid the recovery of native species. To develop or find a biological control that exerts control only on the targeted species is a very lengthy process of research and experiments. Biological pest control 9

Effects on invasive species

Biological control programs aim to reduce or eliminate populations of ecologically and agriculturally harmful invasive species. Examples where this has been achieved include: • The alligator [23] was introduced to the United States from . This aquatic weed spreads rapidly and causes many problems in lakes and rivers. The weed takes root in shallow water causing major problems for navigation, , and . The invasive species Alternanthera philoxeroides The alligator weed beetle and two other biological controls (alligator weed) was controlled in (U.S.) were released in Florida. Because of their success, Florida banned by the introduction of hygrophila the use of to control alligator weed three years after the (alligator weed ) controls were introduced.[24] • calmariensis, a , has been introduced in North America as a control agent for purple loosestrife (). • In the late 19th century cottony cushion scale was discovered in the Californian citrus industry, and it was feared that severe economic losses would result. However the vedalia beetle and, subsequently, Cryptochaetum iceryae, a parasitoid fly, were introduced to control the pest. Within a few years the cottony cushion scale was controlled and the citrus industry suffered little financial loss. • is a major aquatic weed. It covers many waterways causing damage to water flow and the ecosystem. This weed was incorrectly identified at first. Scientists found weevils eating this weed. They named this species of weevils C. salviniae. The weevils have become a great biological control success for all countries. [25]

• Prickly pear cacti, spp., were introduced in to Queensland, Australia as an ornamental plant. It quickly spread to cover over 25 million hectares of Australia. Two control agents were used to help control the spread of the plant. These were C. cactorum, a Lepidopteran species, and spp., a Hemipteran species. [26]

Grower Education A potential obstacle to the adoption of biological pest control measures is growers sticking to the familiar use of pesticides, particularly since it has been claimed that many of the pests that are controlled today using pesticides, actually became pests because use reduced or eliminated natural predators.[27] A method of increasing grower adoption of biocontrol involves letting growers learn by doing, for example showing them simple field experiments, having observations of live predation of pests, or collections of parasitised pests. In the Philippines, early season sprays against leaf folder caterpillars were common practice, but growers were asked to follow a ‘rule of thumb’ of not spraying against leaf folders for the first 30 days after transplanting; participation in this resulted in a reduction of use by 1/3 and a change in grower perception of insecticide use.[28] Biological pest control 10

References

[1] "Biological control: Approaches and Applications" (http:/ / ipmworld. umn. edu/ chapters/ landis. htm). University of Minnesota. . Retrieved 3 September 2012. [2] Follett PA, Duan JJ (2000) 'Nontarget effects of biological control.' (Kluwer Academic Publishers). [3] "the only biogenic amines whose physiological significance is presumably restricted to , pharmacologists have focused their attention on the corresponding receptors, which are still believed to represent promising targets for new " TYRAMINE AND OCTOPAMINE: Ruling Behavior and Metabolism T Roeder - Annual Review of , 2005 - Annual Reviews

[4] Spider mites and their natural enemies (http:/ / 207. 5. 17. 151/ biobest/ en/ films/ dvd. htm)

[5] White flies and their natural enemies (http:/ / 207. 5. 17. 151/ biobest/ en/ films/ dvd. htm) [6] Hoddle M.S., Grandgirard J., Petit J., Roderick G.K., Davies N. (2006). "Glassy-winged sharpshooter Ko'ed - First round - in ". Biocontrol News and Information 27 (3): 47N–62N. [7] Smith SM (1996) Biological control with Trichogramma: advances, successes, and potential of their use. In 'Annual Review of Entomology' pp. 375-406. [8] L.A. Swan. 1964. Beneficial Insects. 1st ed. page 249. [9] I.M. Hall & P.H. Dunn, Entomophthorous Fungi Parasitic on the Spotted Alfalfa Aphid, Hilgardia, Sept 1957.

[10] Gould, Juli; Bauer, Leah. "Biological Control of Emerald Ash Borer (Agrilus planipennis)" (http:/ / www. aphis. usda. gov/ plant_health/

plant_pest_info/ emerald_ash_b/ downloads/ eab-biocontrol. pdf). and Plant Health Inspection Service (APHIS) website (http:/ / www.

aphis. usda. gov/ ) (United States Department of ). . Retrieved 28 April 2011 [11] Bauer, L.S.; Liu, H-P; , D.; Gould, J. (2008). "Developing a classical biological control program for Agrilus planipennis (Coleoptera:

Buprestidae), an invasive ash pest in North America" (http:/ / www. nrs. fs. fed. us/ pubs/ jrnl/ 2008/ nrs_2008_bauer_002. pdf). Newsletter of the Michigan Entomological Society 53 (3&4): 38–39. . Retrieved 29 April 2011.

[12] "Biocontrol: and Wasps Released to Control Emerald Ash Borer" (http:/ / www. sciencedaily. com/ releases/ 2011/ 04/

110426111415. htm). News. ScienceDaily. 26 April 2011. . Retrieved 27 April 2011.

[13] "Factsheet - Mucuna pruriens" (http:/ / www. tropicalforages. info/ key/ Forages/ Media/ Html/ Mucuna_pruriens. htm). www.tropicalforages.info. . Retrieved 2008-05-21. [14] Khan, Z. (2008). "On- evaluation of the ‘push–pull’ technology for the control of stemborers and striga weed on in western Kenya". Field Crops Research 106: 224–233. doi:10.1016/j.fcr.2007.12.002. [15] Bornemissza, G. F. (1976), The Australian dung beetle project 1965-1975, Australian Research Committee Review 30:1-30

[16] http:/ / www. csiropedia. . au/ display/ CSIROpedia/ Dung+ beetle+ program [17] Council NR (1996) 'Ecologically Based Pest Management:New Solutions for a New Century.' (The National Academies Press)

[18] Society for (2002), "Biocontrol backfires again," http:/ / www. scienceblog. com/ community/ older/ 2002/ C/

20025043. html, accessed July 31, 2009. [19] Wright MG, Hoffmann MP, Kuhar TP, Gardner J, Pitcher SA (2005) Evaluating risks of biological control introductions: A probabilistic risk-assessment approach. Biological Control 35, 338-347.

[20] "Cane Toad" (http:/ / www. nt. gov. au/ nreta/ wildlife/ animals/ canetoads/ index. html). Exotic Animals - Major Pests. Northern Territory Government, Australia. . Retrieved 14 March 2011.

[21] http:/ / fdrproject. org. au/ pages/ toads. htm [22] National Research Council (U.S.). Board on Agriculture and Natural Resources (June 2000). Incorporating science, economics, and

sociology in developing sanitary and phytosanitary standards in international trade: proceedings of a conference (http:/ / books. google. com/

books?id=KsQuyCgumREC& pg=PA97). National Academies Press. p. 97. ISBN 978-0-309-07090-4. . Retrieved 12 August 2011.

[23] "Alternanthera philoxeroides information from NPGS/GRIN" (http:/ / www. ars-grin. gov/ cgi-bin/ npgs/ html/ . pl?101502). www.ars-grin.gov. . Retrieved 2008-08-09. [24] Cofrancesco (2007)

[25] http:/ / www. environment. gov. au/ biodiversity/ invasive/ weeds/ publications/ guidelines/ wons/ pubs/ s-molesta. pdf

[26] http:/ / www. daff. qld. gov. au/ documents/ Biosecurity_EnvironmentalPests/ IPA-Prickly-Pear-Control-PP29. pdf [27] Thacker, J.R.M. 2002. An Introduction to Pest Control. Cambridge University Press. [28] Heong, K.L. and Escalada, M. M. 1998, Int. J. Pest Management 44: 191-197 Biological pest control 11

Further reading • Wiedenmann, R. (2000). Introduction to Biological Control. Midwest Institute for Biological Control. Illinois.

Available from http:/ / www. inhs. uiuc. edu/ research/ biocontrol • Cowie R. H. (2001) "Can snails ever be effective and safe biocontrol agents?". International Journal of Pest

Management 47(1): 23-40. PDF (http:/ / www. institutohorus. org. br/ download/ artigos/ Cowie 2001. pdf) • R. James Cook (September 1993). "Making Greater Use of Introduced for Biological Control of

Plant Pathogens" (http:/ / arjournals. annualreviews. org/ doi/ abs/ 10. 1146/ annurev. py. 31. 090193. 000413). Annual Review of Phytopathology 31: 53–80. doi:10.1146/annurev.py.31.090193.000413. PMID 18643761. • U.S. Congress, Office of Technology Assessment (1995). "Biologically based technologies for pest control"

(http:/ / www. princeton. edu/ ~ota/ disk1/ 1995/ 9506/ 9506. PDF) (PDF). OTA-ENV-636 (US Government Printing Office, Washington, DC). • Felix Wäckers, Paul van Rijn and Jan Bruin (2005). Plant-Provided Food for Carnivorous Insects - a protective and its applications. Cambridge University Press, UK, 2005. ISBN 978-0-521-81941-1. Building organic pest-free gardens

• The Time Saving Garden (http:/ / www. readersdigest. co. uk/ timesaving-garden-p-691. html?cPath=41_50) by David and Charles PLC/Reader's Digest, ISBN 978-0-276-44245-2 Effects on native biodiversity • Pereira, M.J. et al. (1998) Conservation of natural vegetation in Azores Islands. Bol. Mus. Munic. Funchal 5, 299–305 • Weeden, C.R., A. M. Shelton, and M. P. Hoffman. Biological Control: A Guide to Natural Enemies in North

America. Available from (http:/ / www. nysaes. cornell. edu/ ent/ biocontrol/ predators/ rodolia_cardinalis. html) (accessed December 2007)

• Cane toad: a case study. 2003. Available from (http:/ / www. biotechnologyonline. gov. au/ enviro/ canetoad. cfm) (accessed December 2007) • Humphrey, J. and Hyatt. 2004. CSIRO Australian Animal Health Laboratory. Biological Control of the Cane Toad Bufo marinus in Australia • Cory, J. and Myers, J. 2000. Direct and indirect ecological effects of biological control. Trends in Ecology & Evolution. 15, 4, 137-139. • Johnson, M. 2000. and Scope of Biological Control. Biological Control of Pests Effects on invasive species • Cofrancesco, A. 2007. U.S. National Management Plan: An Action Plant for the Nation- Control and

Management. Army Corps of Engineers. Available from (http:/ / www. invasivespeciesinfo. gov/ index. shtml) Effects on the future

• Cooksey D. 2002. Biological Control in Pest Management Systems of Plants. (http:/ / nature. berkeley. edu/

bicon/ what is Biological Control. htm) Western Regional Committee, Riverside, . Economic effects • Griffiths, G.J.K. 2007. Efficacy and economics of shelter for conservation. Biological Control: in press. doi:10.1016/j.biocontrol.2007.09.002 • Collier T. and Steenwyka, R. 2003. A critical evaluation of augmentative biological control. Economics of augmentation: 31, 245-256. Biological pest control 12

External links

• Biological Pest Control (http:/ / www. invasivespeciesinfo. gov/ toolkit/ controlmech. shtml), National Invasive Species Information Center, United States National Agricultural Library. Lists general information and resources for biological pest control.

• Biological fight applied to the South countries (http:/ / en. ird. fr/ the-media-library/ videos-on-line-canal-ird/

biological-fight-applied-to-the-south-countries/ lutte-biologique-appliquee-aux-pays-du-sud) - 10 minute video

• Biological Control: A Guide to Natural Enemies in North America (http:/ / www. nysaes. cornell. edu/ ent/

biocontrol/ )

• Construction of a garden and implementation of natural pest control (http:/ / www. ovam. be/ jahia/ Jahia/ pid/ 467)

• Educational movies about biological pest control of white flies, aphids and spider mites in (http:/ /

207. 5. 17. 151/ biobest/ en/ films/ dvd. htm)

• Association of Natural Biocontrol Producers (http:/ / www. anbp. org) Trade association of biological pest control industry

• Alternative Methods of Control (http:/ / entomology. ifas. ufl. edu/ fasulo/ molecrickets/ mcri0001. htm) at the University of Florida News

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